Cl>lAat;i<,<:>';i^;}Oit3Cl <a f^.S^ iVi^ 5' i ''*«0£Mi»«33'**^ SMITHSONIAN INSTITUTION MUSEUM O F NATURAL HISTORY I NotUTus albater, new species, a female paratype, 63 mm. in standard length; UMMZ 102781, Missouri. (Courtesy Museum of Zoology, University of Michigan.) UNITED STATES NATIONAL MUSEUM BULLETIN 282 A Revision of the Catfish Genus Noturus Rafinesque^ With an Analysis of Higher Groups in the Ictaluridae WILLIAM RALPH TAYLOR Associate Curator, Division of Fishes SMITHSONIAN INSTITUTION PRESS CITY OF WASHINGTON 1969 IV Publications of the United States National Museum The scientific publications of the United States National Museum include two series, Proceedings of the United States National Museum and United States National Museum Bulletin. In these series are published original articles and monographs dealing with the collections and work of the Museum and setting forth newly acquired facts in the fields of anthropology, biology, geology, history, and technology. Copies of each publication are distributed to libraries and scientific organizations and to specialists and others interested in the various subjects. The Proceedings, begun in 1878, are intended for the publication, in separate form, of shorter papers. These are gathered in volumes, octavo in size, with the publication date of each paper recorded in the table of contents of the volume. In the Bulletin series, the first of which was issued in 1875, appear longer, separate publications consisting of monographs (occasionally in several parts) and volumes in which are collected works on related subjects. Bulletins are either octavo or quarto in size, depending on the needs of the presentation. Since 1902, papers relating to the botanical collections of the Museum have been published in the Bulletin series under the heading Contributions from the United States National Herbarium. This work forms number 282 of the Bulletin series. Frai^k a. Tatlob Director, United States National Museum U.S. GOVERNMENT PRINTING OFFICE WASHINGTON : 1969 For sale by the Superintendent of Documents, U.S. Government Printing Office Washington, D.C. 20402 - Price $2 Contents Page Introduction 1 Acknowledgments 2 Source of material 4 Methods 5 Counts 5 Measurements 8 The Family Ictaluridae 9 The sensory canal system 11 Relationships of the genera of the Ictaluridae 14 Genus Noturus Rafinesque 20 Key to the subgenera of Noturus 23 History 23 Zoogeography 24 Sexual dimorphism 26 Reproduction 27 Early development 30 Subgenus Schilbeodes Bleeker 32 Key to the species of the subgenus Schilbeodes 34 Noturus gyrinus (Mitchill) 35 Noturus lachneri, new species 54 Noturus exilis Nelson ,. 57 Noturus leptacanthus Jordan 68 Noturus nocturnus Jordan and Gilbert 74 Noturus insignis (Richardson) 83 Thefunehris group 95 Noturus funebris Gilbert and Swain 96 Noturus -phaeus, new species 100 Noturus gilberti Jordan and Evermann 105 Subgenus Noturus Rafinesque 108 Noturus flavus Rafinesque HI Subgenus Rabida Jordan and Evermann 128 Key to the species of the subgenus Rabida 130 The hildebrandi group 131 Noturus hildebrandi (Bailey and Taylor) 131 Noturus hildebrandi hildebrandi (Bailey and Taylor) 136 Noturus hildebrandi lautus, new subspecies 138 Noturus baileyi, new species 141 Noturus albater, new species 144 The elegans group 1^0 Noturus elegans, new species 150 Noturus trautmani, new species 156 Noturus eleutherus Jordan 1^0 The furiosus group 1^^ 168 . Noturus placidus, new species Noturus stigmosus, new species 173 Noturus munitus Suttkus and Taylor 180 V VI CONTENTS Page Noturus furiosus Jordan and Meek 184 The miurus group 188 Noturus miurus Jordan 190 Noturus flavipinnis, new species 201 Noturus flavater, new species 204 Hybridization 209 Noturus gyrinus {MiichWl) X Noturus miurus Jordan 210 Noturus exilis Nelson X Noturus miurus Jordan 214 Phylogeny 216 Literature cited 221 Tables 1-28 261 Plates 1-21 follows 307 Index 311 A Revision of the Catfish Genus Noturus Rafinesque, With an Analysis of Higher Groups in the Ictaluridae Introduction This study of the small North American catfishes in the genus Noturus was undertaken to analyze the species and to determine their relationships. It is based on critical examination of most of the many thousand specimens of Noturus now in museums, upon several hundred skeletonized or cleared and stained specimens of catfishes, and upon comparison with most of the other species in the family Ictaluridae. The species in the family Ictaluridae appear to constitute six genera, forming three major natural groups. Each group includes one mono- typic genus that is blind, unpigmented, and of restricted subterranean range, and one genus with species that are eyed, pigmented, and of widespread distribution in surface waters. The divisions are: (1) an Ictalurus group including the genus Ictalurus Rafinesque and the blind Trogloglanis pattersoni Eigenmann, (2) a Noturus group containing that genus and Prietella phreatophila Carranza, and (3) the Pylodictis group consisting of the large Pylodictis olivaris (Rafinesque) and the eyeless Satan eurystomus Hubbs and Bailey. The 23 species of Noturus include one subspecies and ten species that are described here as new. They are arranged in three subgenera: Rabida Jordan and Evermann, Noturus Rafinesque, and Schilbeodes Bleeker. With Prietella, several of their characters are intermediate between those of the Ictalurus group and those of the Pylodictis group. In certain characters they appear to be the most primitive, but they have specialized in several directions, exemplified by loss, reduction, or fusion of some structures and by increase in others. Hubbs and Raney (1944) studied much of the available material of Noturus exilis, Noturus insignis, and Noturus gUberti, giving some characters and mapping distributions. However, they incorrectly changed the names of exilis, insignis, and gyrinus to insignis, margin- atum, and mollis, respectively, in the then recognized genus Schilbeodes. Otherwise, no study of the genus Noturus utilizing the existing material has been made since Jordan (1877d) and Swain and Kalb (1883). The species of Noturus are mainly active at night, hiding in cavities or beneath objects during daylight. Consequently, they are often most readily collected by using chemicals, direct current electricity, or by seining after dusk. Some of the species appear to be very spottily 1 2 U.S. NATIONAL MUSEUM BULLETIN 282 distributed and are thus infrequently obtained except in intensive survey work. One, Noturus flavipinnis has not been collected since 1893. The known localities from which it was obtained are now ecologically unsuitable for its existence and it may be extinct. Although not often encountered, the members of the genus become rather widely known because of the painful "sting" produced by their pectoral spines. The study of the variation and distribution of populations in Noturus has revealed several interesting patterns and presented problems of interpretation. Some wide ranging species, particularly miurus and insignis, show relatively little structural change geographically, but several structures in Noturus gyrinus gradually increase in length and number from north to south. Similarly a west to east increase is ap- parent in pectoral rays of Noturus leptacanthus and in anal rays of Noturus funebris. The presence of highly variant populations of Noturus nocturnus in the Red River system, which is in the middle of its range, is puzzling. In a number of species, localized and relatively minor variation is noted. Three clusters of species, each with closely related allopatric repre- sentatives, are recognized as species groups; other species groups are suggested but the relationship of the species is not as clear. These allopatric populations are recognized as species because they show a relative morphological uniformity, with consistent divergence from related populations; trends or gradation of characters from one popula- tion toward the other do not exist and intermediates or intergrades are lacking. Natural hybrids between species of Noturus are described but hy- bridization is rare. Sexual dimorphism in the family, once thought to be important in the interpretation of species of Noturus, has been exaggerated ; instead differential morphological change, accompanying growth and sexual maturity, has resulted in several unnecessary names in Ictalurus, complicating its nomenclature. Acknowledgments A great many individuals have contributed importantly to this study in many ways, especially by the loan of specimens or making them available for study, provision of laboratory space, aid in col- lecting specimens, or have given freely information about specimens, notes on specimens in their care, or data from writings unavailable to me. To each I am greatly indebted and I sincerely trust that no one has been overlooked: Reeve M. Bailey, Marie-Louise Bauchot, Norman Benson, the late Leon Bertin, Albert P. Blair, James E. Bohlke, E. Milby Burton, Robert S. Campbell, Jorge Carranza, William M. Clay, Robert E. Cleary, Bruce B. Collette, the late CATFISH GENUS NOTURUS RATINESQUE 6 Fannye A. Cook, Walter R. Courtenay, Jr., Frank B. Cross, Myvanwy M. Dick, Samuel Eddy, D. H. Enlow, Charles E. Farrell, the late Henry W. Fowler, Barry 0. Freeman, Glenn Gentry, Shelby D. Gerking, Carter Gilbert, Francois Gouin, P. H. Greenwood, Harold W. Harry, Clark Hubbs, Stanton Hudson, Robert H. Kanazawa, Leslie W. Knapp, Ernest A. Lachner, Robert E. Lennon, L. W. Lowe, Y. J. McGaha, Giles W. Mead, Robert R. Miller, George